PHYLOGENETIC IMPLICATIONS OF 
GROOMING BEHAVIOR IN COCKROACHES 
(INSECTA: BLATTARIA)* 
By Bonnie J. Bobula Smith and Barry D. Valentine 
Department of Zoology, The Ohio State University, 
Columbus, Ohio 43210-1293 
Introduction 
The use of behavioral data for the investigation and delineation of 
evolutionary relationships has been increasingly heuristic (Evans, 
1952; Speith, 1952; Alexander, 1962; etc.). In insects, grooming 
behavior seems particularly useful because it is a widespread and 
prominent part of their behavioral repertory. Comparative studies 
of several orders have been based on grooming behavior (Szy- 
manski, 1918; Heinz, 1949; Gangwere, 1958; Jander, 1966; Farish, 
1972; Valentine, 1973; Valentine and Glorioso, 1979). Lipps (1973) 
provides an excellent review of grooming literature. 
The cockroaches are a diverse group with an extensive nontaxo- 
nomic literature. There are many descriptions of the general biology 
of the order (Gould and Deay, 1938; Roth and Willis, 1954, etc.) 
and of particular species (Qadri, 1938; Rau, 1940; Dow, 1955; Hes- 
lop and Ray, 1959, etc.). Roth and Willis (1954) and McKittrick 
(1964) summarize many such papers, especially on biology. Our 
taxonomy follows McKittrick (1964), and is summarized in Table 1. 
Unspecified grooming behavior of cockroaches is mentioned by 
Patton (1941) and Burkholder (1965). Others discuss grooming of 
particular body parts (Haber, 1920; Mote, Wilcox, and Davis, 1926; 
Roth and Willis, 1952 and 1954; and Eisner, 1961). A few authors 
are more detailed: Gangwere (1958), Yoshikawa (1958), Eaton and 
Farley (1969), and Lipps (1973). The most complete description of 
roach grooming is by Turner (1913), who describes in excellent 
detail antennal, palpal, and leg cleaning using the mouth, and the 
use of a leg to rub the base of the antenna and dorsal surface of the 
abdomen, all in a roach he called Periplaneta orientalis, now placed 
in the genus Blatta. 
* Manuscript received by the editor July 15, 1985. 
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